High-temperature tolerance of photosynthesis in the red alga Chondrus crispus

Chondrus crispus (Stackhouse) is a perennial red seaweed, common in intertidal and shallow sublittoral communities throughout the North Atlantic Ocean. In the intertidal zone, C. crispus may experience rapid temperature changes of 10 to 20C° during a single immerison-emerision cycle, and may be exposed to temperatures that exceed the thermal limits for long-term survival. C. crispus collected year-round at Long Cove Point, Chamberlain, Maine, USA, during 1989 and 1990, underwent phenotypic acclimation to growth temperature in the laboratory. This phenotypic acclimation enhanced its ability to withstand brief exposure to extreme temperature. Plants grown at summer seawater temperature (20°C) were able to maintain constant rates of lightsaturated photosynthesis at 30°C for 9 h. In contrast, light-saturated photosynthetic rates of plants grown at winter seawater temperature (5°C) declined rapidly following exposure to 30°C, reached 20 to 25% of initial values within 10 min, and then remained constant at this level for 9 h. The degree of inhibition of photosynthesis at 30°C was also dependent upon light intensity. Inhibition was greatest in plants exposed to 30°C in darkness or high light (600 mol photons m^-2s^-1) than in plants maintained under moderate light levels (70 to 100 mol photons m^-2s^-1). Photosynthesis of 20°C-acclimated plants was inhibited by exposure to 30°C in darkness or high light, but the degree of inhibition was less than that exhibited by 5°C-grown plants. Not only was light-saturated photosynthesis of 20°C plants less severely inhibited by exposure to 30°C than that of 5°C plants, but the former also recovered faster when they were returned to growth conditions. The mechanistic basis of this acclimation to growth temperature is not clear. Our results indicate that there were no differences between 5 and 20°C-grown plants in the thermal stability of respiration, electron transport associated with Photosystems I or II, Rubisco or energy transfer between the phycobilisomes and Photosystem II. Overall, our results suggest that phenotypic acclimation to seawater temperature allows plants to tolerate higher temperatures, and may play an important role in the success of C. crispus in the intertidal environment.     

Autecology and clonal variability of the marine centric diatom Thalassiosira rotula (Bacillariophyceae) in response to light,temperature and salinity

The influence of 49 combinations of salinity (10–40 S, at 5 S intervals) and temperature (0°–30°C, at 5C° intervals) on the maximum daily division rate (K) and 18 combinations of light intensity (six levels) and temperature (5°, 15°, and 25°C) on photosynthesis, cell division, and chlorophyll a was examined using two clones of Thalassiosira rotula Meunier isolated from the upwelling area of Baja California (clone C8) and from Narragansett Bay, Rhode Islands (clone A8). Physiological differences appear to characterize these to clones with regard to their temperature tolerance (C8 5°–30°C, A8 0°–25°C), maximum growth rate (C8 K=2.9, A8 K=2.4), chlorophyll a content, and in the rates of growth and photosynthesis in response to light intensity and temperature. Optimum salinity for both clones (25–30 S) was generally independent of temperature, while chlorophyll a content decreased with temperature. T. rotula is a cosmopolitan paractic species; experimental studies indicate that it is eurythermal and moderately euryhaline. Comparison of five additional Narragansett Bay isolates of T. rotula reveal minimal spacial or temporal variability in genetically determined physiological characteristics within this local population.     

Photosynthesis and amelioration of desiccation in the intertidal saccate alga Colpomenia peregrina

Rates of gross photosynthesis for the intertidal saccate alga Colpomenia peregrina (Sauv.) Hamel were determined under submersed and emersed conditions. Maximal photosynthetic rates were lower than for most seaweeds but comparable with other saccate members of the genus. By fitting the data to a hyperbolic tangent function, maximal photosynthetic rates were estimated to be 5.29 mmol CO₂ m^-2 h^-1 under submersed conditions and 2.06 mmol CO₂ m^-2 h^-1 under emersed conditions. I_k for submersed thalli was 69.1 E m^-2 s^-1, wherea for emersed thalli it was 149.0 E m^-2 s^-1, or 2.2 times higher. At low tide in the field and under saturating irradiance, carbon from seawater retained within the thallus cavity was assimilated at 0.9 mmol CO₂ m^-2 h^-1. In the laboratory under emersed conditions, carbon from this source was taken up at 0.6 mmol CO₂ m^-2 h^-1 at 20°C and at 0.34 mmol CO₂ m^-2 h^-1. Retained seawater also greatly reduced drying under desiccating conditions. Experimental thalli from which seawater had been removed lost thallus water continuously throughout the drying period (120 min). On the other hand, control, thalli lost water for the first 15 min, after which no further water loss occurred. At the termination of the experiment, control thalli had lost 7.2% of their water, whereas experimental thalli had lost 39.2%. Desiccation affected the emersed photosynthetic rate of experimental and control thalli. Emersed photosynthetic rates for thalli dried for 15 min were higher than for fullyhydrated thalli. However, emersed photosynthesis of thalli dried for longer than 15 min was lower than fully-hydrated rates and was directly related to percent water loss. Utilizing data from this study, a model was constructed to determine total photosynthetic production of C. peregrina over a single daylight period. From these calculations it was determined that emersed photosynthesis can increase daily photosynthetic production of C. peregrina by 50%.     

Effect of freezing on photosynthesis of intertidal macroalgae: relative tolerance of Chondrus crispus and Mastocarpus stellatus (Rhodophyta)

The effect of freezing on photosynthetic metabolism was studied in the red algae, Chondrus crispus and Mastocarpus stellatus. Plants of both species were collected from the intertidal at Chamberlain or Kresge Point, Maine, USA (43°56N, 69°54W) between February and March 1987. Photosynthetic rates were measured immediately after freezing at-20°C and following recovery periods in seawater. Photosynthesis in C. crispus declined rapidly following freezing, falling to 70% of control values within 1 h and 30% after 3 h exposure. Minimum photosynthetic rates (7 to 9% of controls) occurred following freezing exposures of 12 h or more. Full photosynthetic recovery in C. crispus after 3 h at-20°C required 48 h. Photosynthesis in C. crispus did not fully recover in plants frozen for 6 h or more. In contrast, photosynthesis in M. stellatus was relatively unaffected by freezing exposures of <12 h. Twelve hours or more at-20°C reduced photosynthesis to 55% of controls. Photosynthesis in M. stellatus fully recovered from 24 h at-20°C within 24 h. In both species the reduction of photosynthesis by freezing was associated with damage to the plasma membrane and reduced efficiency of energy transfer from phycobilisomes to chlorophyll a, but did not appear to involve ribulose-1,5-bisphosphate carboxylase oxygenase activity. The freezing tolerance of C. crispus and M. stellatus positively correlates with their respective intertidal distributions, suggesting that freezing may be involved in controlling the distributions of these species on the shore.     

Influence of high temperature and residual chlorine on marine phytoplankton

Marine phytoplankton forms are frequently exposed to sudden biological changes such as rapid rise in water temperature and chlorine content of their environment, resulting from the use of sea water for cooling purposes by electric generators. The direct influence of these effluents, i.e. inhibitory effects of high temperature and residual chlorine on growth and photosynthesis of Chlamydomonas sp. and Skeletonema costatum, were investigated experimentally. Chlamydomonas sp. and S. costatum exposed to high temperatures were affected in their growth from 43° and 35°C, respectively, by immersion of the respective cultures in a warm bath for 10 min. Treatment at high temperatures of 40 °C and 30° 35°C for 10 min, influenced their photosynthetic activities, which were completely inhibited immediately after 10 min exposure at 42° and 37 °C, respectively. S. costatum was killed by chlorine at a concentration of 1.5 2.3 ppm when exposed for exactly 5 or 10 min, while Chlamydomonas sp. was not irreversibly damaged even at 20 ppm chlorine or more with the same exposure period. These results lead to the conclusion that the high temperature of, and residual chlorine in, effuents from a power plant discharging into the open sea, should not cause great damage to marine phytoplankton in that area.     

Interactions between light and temperature on the physiological ecology ofGracilaria tikvahiae (Gigartinales: Rhodophyta)

Main effects and interactions of light and temperature on rates of growth (), net photosynthesis (P_s), and dark respiration (R) of the red seaweedGracilaria tikvahiae were investigated in outdoor, nutrient-replete continuous-flow seawater culture chambers. Below 15°C,G. tikvahiae did not grow and between 15° and 30°C, both main effects and interactions of light and temperature on and P_s were significant, which explains the occurrence of this alga as a summer annual in its northern range. Temperature interacted with light (I) through its influence on the vs I and P_s vs I curves. The initial slope of the vs I curve, , the light saturation intensity, I_s, and maximum growth rate, _max, were all significantly lowerat 15°C compared to 20°, 25°, or 30°C. Maximum values of _max, the P_s:R ratio and the net photosynthesis:gross photosynthesis ratio (P_s:P_g) all occurred at 25°C, suggesting that this is the best temperature for growth ofG. tikvahiae. Values for P_max increased up to 30°C, indicating that the temperature for maximum growth and net photosynthesis are not the same forG. tikvahiae. Significant photoinhibition of growth and photosynthesis at full incident sunlight (I₀) occurred at 15°C but not at 20°, 25°, or 30°C. Steele's equation fit the 15°C vs I data best, whereas the hyperbolic tangent function fit the 20°, 25°, and 30°C data best. Main effects and interactionof light intensity and temperature on rates of R were also significant (P<0.001). R was highly intercorrelated with and P_s (0.86r0.94), indicating that R inG. tikvahiae is primarily regulated by growth rate and not temperatureper se. Environmental factors that regulate growth, such as light intensity, exert a great influence on R inG. tikvahiae.     

Temperature ecotypes near the southern boundary of the kelp Laminaria saccharina

Effects of temperature on survival, growth, and photosynthesis were compared for two USA populations of Laminaria saccharina Lamour. One population was located in New York State, near the southern latitudinal boundary of the species in the western North Atlantic. This southern boundary population was exposed to ambient temperatures 20°C for about 6 wk each summer. The second population was located in Maine, toward the center of the latitudinal range of the species, and was rarely exposed to temperatures>17°C. sporophytes from the New York (NY) population exhibited greater tolerance of high temperature than plants from the Maine (ME) site. Juvenile sporophytes from the two sites had similar rates of survivorship and growth at temperatures below 20°C, but showed different responses at 20°C in laboratory experiments. NY plants survived and grew for 6 wk at 20°C. ME plants showed negative growth during wk 2 and 100% mortality during wk 3. NY and ME plants held in situ at the NY site during June to September, 1985, also exhibited differential survivorship when ambient temperatures exceeded 20°C. Results of photosynthesis and dark respiration measurements on NY and ME plants grown at various temperatures suggested that the high-temperature tolerance of NY plants was attributable to their ability to maintain positive daily net C-fixation at 20°C. The high-temperature tolerance of the NY plants appeared to be due to genetic adaptation and is probably crucial to the persistence of the species near its southern boundary.     

Physiological responses of Mytilus edulis and Cardium edule to aerial exposure

Physiological responses of two bivalves (Mytilus edulis L. and Cardium edule L.) to intertidal conditions were studied. Specimens were collected from S. W. England in autumn/winter, 1980, and acclimatized to either intertidal or subtidal regimes before measurement of rates of heat dissipation and oxygen uptake during 5 h of air exposure, and rates of , particle clearance, ammonia excretion, and food-absorption efficiency during 7 h of reimmersion. Subtidal individuals were either intermittently or continuously fed in order to distinguish the effects of periodic food supply from the effects of air exposure. Specimens of M. edulis had low aerial rates of (14 to 20% of aquatic rate), and was greater than the energy equivalent of , indicating that they were largely anaerobic. In contrast, C. edule air-gaped and had higher aerial rates of and (50 to 75% of aquatic rate). There were behavioural and metabolic differences in the responses of intertidally and subtidally acclimatized mussels and cockles to air exposure. Intertidal individuals of both species were more quiescent, had lower aerial rates of and , and showed a conditioned response at the expected time of reimmersion. The reduction in aerial rate of was an energy-saving mechanism and the payment of the oxygen-debt within 2 h of reimmersion represented a significant cost. The heat increment associated with feeding and digestion was estimated as 15 to 17% of the oxygen uptake by M. edulis during all stages of recovery. M. edulis adapted to the intertidal regime by reducing its time-averaged aerial and aquatic rates of ammonia excretion. In contrast, C. edule maintained a high aerial and aquatic rate of ammonia excretion. The clearance rates of M. edulis recovered rapidly (0.5 to 1.5 h) following reimmersion, whereas those of C. edule recovered more slowly, particularly for the subtidal individuals following acute exposure (>4 h). There was no evidence of increased clearance rate or absorption efficiency by intertidal individuals to compensate for the loss of feeding time. Intertidally acclimatized individuals of M. edulis and C. edule had more energy available for growth (scope for growth) integrated over a 12 h period and higher growth efficiencies than subtidal individuals subjected to acute air exposure and intermittent feeding regimes. This was the result of reduced aerial and aquatic rates of energy expenditure, a relatively small cost in terms of the payment of oxygen-debt and a rapid recovery of clearance rate following reimmersion.     

Improved survival under heat stress in intertidal embryos (<Emphasis Type="Italic">Fucus</Emphasis> spp.) simultaneously exposed to hypersalinity and the effect of parental thermal history

Intertidal organisms exposed to thermal stress normally experience other stresses simultaneously, but how these combined stresses modify tolerance to heat, especially for embryos, is poorly understood. Tolerance of fucoid algal embryos to heat, with and without acclimation to a sublethal temperature and with simultaneous exposure to hypersaline media, was examined. Embryos of Fucus vesiculosus L. (mid-intertidal zone) were less tolerant than embryos of Fucus spiralis L. (upper intertidal zone); without acclimation and with a growth temperature of 14°C, about half of the embryos survived 3 h exposure to 33°C in F. vesiculosus and of 35°C in F. spiralis. Conditions experienced by parental thalli (4°C versus 14°C storage) significantly affected the heat tolerance of embryos grown for 24 h post-fertilization at 14°C in F. vesiculosus, a result that is important for biologists using fucoid algae as model systems. Acclimation to a sublethal temperature (29°C) or exposure to the LT₅₀ (33°C, F. vesiculosus; 35°C, F. spiralis) in 100 psu seawater (2850 mmol kg^–1 osmolality) resulted in 30–50% higher levels of embryonic survival. Higher levels of HSP60s were found in embryos exposed to 29–33°C than to 14°C; lower levels of HSP60s were present in embryos exposed to the LT₅₀ under hypersaline conditions than in normal seawater. Contemporaneous studies in 1995–1996 of substratum temperature and desiccation levels were made at Schoodic Point, Maine (USA) underneath F. spiralis and F. vesiculosus canopies and in Semibalanus balanoides patches. This study extends the bioindicator utility of heat-shock proteins in studies of intertidal organisms and demonstrates the importance of integrated stress responses in survival of a single stress factor (e.g. temperature).Electronic Supplementary Material Supplementary material is available in the online version of this article at     

The significance of temperature,salinity and zinc as lethal factors for the mussel Mytilus edulis in a polluted estuary

Mussels, Mytilus edulis L., were subjected to high temperatures, low salinities and dissolved zinc in order to investigate possible environmental hazards of a discharge of heated effluent near Newport on the Yarra River estuary, Victoria, Australia. Exposure to zinc at 0.8 mg l^-1 for 14 d in otherwise favourable conditions significantly increased mortality resulting from subsequent exposure to temperatures between 29° to 31°C for 24 h without added zinc. Mussels collected from water of temporarily lowered salinity (8–16 S) showed significantly lower thermal resistance than controls collected from marine salinities (35 S). Mussels taken from a marine environment and exposed to 10 S died at a rate which increased with temperature. Mussels acclimated for 14 d to combinations of 10°, 16° and 22°C and 22 and 35 S, and subsequently exposed to increased zinc concentrations accumulated zinc to levels which were independent of temperature and salinity. The zinc was lethal more quickly at 22°C and 35 S than at the lower temperatures and salinities. The modes of toxic action of the salinity, zinc and temperature factors are discussed and it is argued that zinc which has been found accumulated in mussels near Newport could be reducing their resistance to raised temperatures and perhaps other stresses, probably as a result of effects on lysosomal functioning. The evidence suggests that the heated effluent will accelerate any toxic effects of zinc or low salinities which occur near Newport and so poses a hazard in winter as well as in summer.     

Seasonal effects of light,temperature, nutrient concentration and salinity on the physiology and growth of Caulerpa paspaloides (Chlorophyceae)

Caulerpa paspaloides (Bory) Greville were collected during the winter and summer (1978 to 1979) from the Florida Keys, USA. Thalli collected during the winter photosynthesized more efficiently at low light intensities (I_c<1, I_k=38 Exm^-2xs^-1) than did thalli collected in the summer (I_c=13, I_k=111 Exm^-2xs^-1). Summer thalli exhibited higher P_max values (2.20 mgO₂xg^-1 dry wtxh^-1) than winter thalli (1.70 mg O₂xg^-1 dry wtxh^-1). Rates of rhizome elongation and frond initiation were strongly inhibited by winter temperatures. The maximum lethal temperature for summer thalli was 37° to 38°C as measured by both growth and photosynthesis. No evidence of nitrogen or phosphorus limitation was found. Relatively minor reductions in salinity (3S) resulted in significant increases in rhizome apex motality. Results indicate that low winter temperatures are responsible for reduced winter growth rates previously reported for the Key Largo population. Increased photosynthetic efficiency at low light intensities and preferential maintenance of rhizome elongation over frond initiation appear to allow this tropical macroalga to optimize growth and survival during the winter.     

Temperature acclimation of respiration and photosynthesis in the brown algaLaminaria saccharina

Sporophytes of the brown algaLaminaria saccharina (L.) Lamour grown at 15°C contained significantly more chlorophylla (chla) than did similar plants grown at 5°C. The increase in chla in 15°C plants was due to increased numbers of photosystem II reaction centes, and possibly to increased photosynthetic unit size, compared with 5°C plants. These changes were associated with increased values (photosynthetic efficiencies) in 15°C-grownL. saccharina relative to 5°C-grown plants. The changes in together with reduced respiration rates allowed 15°C-grownL. saccharina to achieve net photosynthesis and light-saturated photosynthesis at a lower photon fluence rate (PFR) than 5°C plants when both groups were assayed at the same temperature (15°C). The photon fluence rates necessary to reach the compensation point and achieve light-saturated photosynthesis (I _c andI _k , respectively) increased with increasing incubation temperature inL. saccharina grown at both 5 and 15°C. However, acclimation responses to growth temperature compensated for the short-term effect of temperature onI _c andI _k . Consequently, plants grown at 5 and 15°C were able to achieve similar rates of light-limited photosynthesis, and similarI _c andI _k values at their respective growth temperatures. These responses are undoubtedly important for perennial seaweeds such asL. saccharina, which frequently grow in light-limited habitats and experience pronounced seasonal changes in water temperature.Please address all correspondence and requests for reprints to I.R. Davison     

Effect of phosphate and ammonium levels on photosynthetic and respiratory responses of the red alga Gracilaria verrucosa

Gracilaria verrucosa (Hudson) Papenfuss exposed to nutrient enriched media (0.1 mM PO₄; 1.0 mM NH ₄ ⁺ ) by pulse feeding 2 h every third day for a period of 5 wk at 20°C and 25–30 salinity showed significantly higher rates of photosynthesis regardless of photon flux density correlated with increased pigment levels. Algae in nonenriched media showed significantly higher levels of soluble carbohydrates and decreased levels of phycoerythrin and chlorophyll a. Photosynthetic and respiratory responses to temperature 15°, 25°, 30°C and salinity (15, 25, 30 S) combinations indicate broad tolerances by both nutrient enriched and non-nutrient enriched algae. Photosynthetic and respiratory rates were highest at the high temperatures. Pulse-fed algae had significantly higher photosynthetic rates than non-nutrient enriched plants at all temperature and salinity combinations. Non-nutrient enriched algae had significantly higher respiratory rates than nutrient enriched algae at only 30°C and 15. The respiratory rates of both nutrient enriched and non-nutrient algae decreased under combinations of higher temperatures and salinities. G. verrucosa, grown without nutrients, has lower tolerances to environmental stresses.     

Foraging strategies of the marine iguana,Amblyrhynchus cristatus

Summary Two foraging strategies were found in marine iguanas (Amblyrhynchus cristatus); (1) subtidal feeding: the animals swam out to sea and dived for algae in the subtidal zone; (2) intertidal feeding: the animals foraged around low tide in the intertidal zone on more or less exposed algae. Most marine iguanas were very consistent in their foraging strategy and so could be classified as subtidal feeders (SFs) or intertidal feeders (IFs). Feeding strategy was weight-related (Fig. 1), not sexspecific. Animals 1,200 g were IFs, animals >1,800 g SFs. Some iguanas in between followed a mixed foraging strategy. SFs foraged independently of the tides, IFs always around low tide (Figs. 2, 3). Feeding time patterns of IFs and SFs are described (Table 1). Sea motion seemed to have little effect on the foraging pattern of SFs, but strongly influenced that of IFs (Fig. 2). The smaller a marine iguana, the faster it cooled when immersed in water (Fig. 4). The difference between water temperature and core temperature of animals returning from foraging was significantly greater in IFs than SFs (Fig. 5). SFs swimming in very cold water regulated their body temperature to prevent excessive cooling. Possible costs and benefits of the two foraging strategies are discussed. Only part of a marine iguana population lives really amphibiously and only ca. 5% of a 24 h day is spent close to or in the water. All social activities, including mating, take place on land. These life history characteristics preclude those adaptations to an amphibious way of life that would at the same time reduce the iguanas' ability to be maximally active at their typical terrestrial body temperature of 35° C.     

Photoacclimation of Ulva rotundata (Chlorophyta) under natural irradiance

Two vegetative clones (designated 11/85 and 7/86 in accordance with month/year of collection) of the green macroalga Ulva rotundata were collected in the vicinity of Beaufort, North Carolina, USA. Each was grown in an outdoor continuous-flow system in summer (20°C) of 1986 and late winter (10° to 17°C) of 1987, in irradiances ranging from 9 to 100% of full sunlight, with and without NH ₄ ⁺ enrichment. Continuous enrichment of influent estuarine water (dissolved inorganic nitrogen 2 M, N:P5) to 8–12 M NH ₄ ⁺ had only a slight effect on growth rate. Temperature changes of 2 to 3°C had a much greater effect. Prolonged exposure to a given daily irradiance resulted in acclimation, exposure to a given daily irradiance resulted in acclimation, indicated by faster growth of conditioned plants relative to those transferred from a different irradiance. Most of the difference in growth rates between transferred and control plants was attributed to differences in thallus absorptance. Growth was photoinhibited above 40% sunlight at temperatures below 15°C, but not above 20°C. Following interday irradiance transfers, thallus percent dry weight changed in a manner that suggests different response times for photosynthesis and cell division.     

Testing the effects of wave exposure,site, and behavior on intertidal mussel body temperatures: applications and limits of temperature logger design

The rocky intertidal mussel Mytilus californianus is exposed to potentially damaging thermal conditions during low tide. However, because the temperatures of ectothermic organisms are driven by multiple climatic factors, we do not fully understand what the body temperatures of intertidal invertebrates are under field conditions, or how thermal stress varies between intertidal sites. We designed a temperature logger that thermally matches (similar size, color, morphology and thermal inertia) living mussels of the species M. californianus, and tested its ability to provide realistic measurements of body temperature in the field. As part of these tests, we examined the propensity of M. californianus to gape, a behavior in which the mussel opens its shell valves, and which may permit evaporative cooling. Because our instruments were unable to mimic this behavior, we tested the degree to which gaping contributes to animal cooling by exposing M. californianus to a range of climatic conditions while recording body temperatures, gaping behavior and water loss. Results indicated no significant influence of gaping on body temperature. Tests comparing temperatures of loggers to those of real mussels under laboratory and field conditions showed that thermally matched loggers recorded temperatures within a few degrees of living animals and that unmodified loggers regularly incurred errors of up to 14°C. We then deployed a series of thermally matched loggers at two sites in central Oregon (Boiler Bay and Strawberry Hill) previously hypothesized to display site-specific differences in aerial temperature, and at various wave-exposure regimes within each site. Significant differences were demonstrated between sites using a subset of temperature metrics in a multivariate analysis. Yearly peaks in maximum temperature, average daily maximum temperature, and degree hours were useful in discriminating between sites. In 2001, wave-exposed sites at Strawberry Hill displayed fewer degree–hours than wave-protected sites, but an equivalent or greater maximum temperature. In 2002 both of these metrics were significantly lower in wave-exposed sites. Boiler Bay and Strawberry Hill differed in thermal regime, but not in a way that would indicate one was hotter than the other.Communicated by P.W. Sammarco, Chauvin     

Effect of temperature on the egg and yolk-sac larval development of common pandora,<Emphasis Type="Italic"> Pagellus erythrinus</Emphasis>

Temperature is one of the most critical environmental factors for fish ontogeny, affecting the developmental rate, survival and phenotypic plasticity in both a species- and stage-specific way. In the present paper we studied the egg and yolk-sac larval development of Pagellus erythrinus under different water temperature conditions, 15°C, 18°C and 21°C for the egg stage and 16°C, 18°C and 21°C for the yolk-sac larval stage. The temperature-independent thermal sum of development was estimated as 555.6 degree-hours above the threshold temperature (the temperature below which development is arrested), i.e. 7°C for the egg and 12.1°C for the yolk-sac larval stage. Higher hatching and survival rates occurred at 18–21°C. At the end of the yolk-sac larval stage, body morphometry differed significantly (p<0.05) between the temperatures tested. The growth rate of the total length increased as temperature rose from 16°C to 18°C, while in the range of 18–21°C it stabilized and was independent of water temperature. The estimated Gompertz growth curve for the yolk-sac larvae of P. erythrinus was (r²=0.992) for the 16°C, (r²=0.991) for the 18°C and (r²=0.981) for the 21°C treatment. The efficiency of vitelline utilization during the yolk-sac larval stage was higher at 18°C.Communicated by O. Kinne, Oldendorf/Luhe     

Adaptations of Gracilaria pacifica (Rhodophyta) to nitrogen procurement at different intertidal locations

Nitrogen uptake, assimilation and accumulation were studied in three populations of Gracilaria pacifica Abbott in Bamfield Inlet, British Columbia, Canada, over three summers, 1979–1981. Two of these populations were in the intertidal one high and one low, and the third was a subtidal cultured population. Nitrate uptake rates, internal nitrate content and nitrate reductase activities were highest in the low intertidal population. Time-courses of uptake and uptake kinetics were studied. Both nitrate and ammonium were taken up simultaneously. Thalli from the high-intertidal population showed enhanced nitrate and ammonium uptake following mild desiccation, and greater tolerance to desiccation in terms of maintaining nitrogen uptake after severe desiccation. Transplants were made to determine the effect of intertidal height and geographic location on responses to desiccation, nitrogen uptake, assimilation and accumulation. Nitrate and ammonium uptake rates were dependent on intertidal height and geographic location. Transplanting up the intertidal increased nitrate uptake and nitrate reductase activity, but decreased the nitrate content of the thalli. There were few significant differences in ammonium uptake rates, and ammonium, amino acid, and soluble-protein content of the various populations. All high-intertidal populations, transplanted or natural, showed enhanced nitrate uptake rates following desiccation. Enhanced ammonium uptake rates following desiccation were restricted to the high-intertidal thalli in only one geographic location. Tolerance to higher levels of desiccation also appeared to be intertidal height-dependent, but required more than five weeks to fully develop or disappear.     

Measuring the effects of thermoclines on the vertical migration of larvae of Callinectes sapidus (Brachyura: Portunidae) in the laboratory

An experimental system is described that is capable of maintaining precise and stable thermoclines of specified magnitude for laboratory study of behavior of planktonic organisms. Behavioral responses to encounters with thermoclines of various magnitudes were measured in stage I larvae of the portunid crab Callinectes sapidus Rathbun. Larvae adapted to an initial temperature of 15°C were able to swim up through a temperature gradient of 10°C, but showed increasing inhibition of upward migration when t=12.5°C. Larvae adapted to 17.5°C were unable to penetrate t=10°C. Thermoclines of 5° or 10°C slowed passive sinking rates. Results suggest that larvae avoid an upper temperature of 27.5°C by a cessation of swimming, accompanied by passive sinking. Thermal gradients of the magnitude present in the natural habitat are unlikely to alter larval vertical migration that is regulated by other exogenous stimuli. Naturally occurring thermoclines should seldom influence dispersal characteristics of C. sapidus larvae.Contribution No. 1535 of the Center for Environmental and Estuarine Studies, University of Maryland     

Mechanisms to initiate a Heterosigma akashiwo red tide in Osaka bay

The ecological role of diel vertical migration of Heterosigma akashiwo Hada to initiate red tide was investigated in Tanigawa Fishing Port and Sano Harbor, Osaka Bay, Japan during red tide seasons in 1979 and 1980. This species migrated toward the surface early in the morning at a velocity of 1.0 to 1.3 m h^-1. Downward migration was found in the afternoon, and more than 2.0×10³ cells ml^-1 aggregated in the bottom layer at night. The upward migration started before sunrise and downward shifting occurred prior to sunset. The movement is presumably associated with circadian rhythm, which is known as one of the biological periodicities. H. akashiwo crossed steep temperature and salinity gradients (6.5°C and 5.7 S) during the diel vertical migration. High values of particulate organic carbon and nitrogen concentrations were obtained in dialysis bags suspended in situ at identical layers with high cell concentration, while the values for surface and bottom bags were comparatively low. The results reveal that H. akashiwo migrates toward the sea surface to carry out photosynthesis effectively, and to the bottom to utilize nutrients efficiently.